BULLETIN OF MARINE SCIENCE.

30(4):858-870, 1980

A CLARIFICATION OF THE GENUS PERNA (MYTILIDAE)

Scott E. Siddull

ABSTRACT
Many morphological features of adult mussels are variable and often of limited value in
establishing taxonomic relationships. Synonymies in the Mytilidae are numerous, the genus
Perna notwithstanding. The historical development and geographical distribution of the three
species placed in the genus Pema are reviewed.
As a post-metamorphic character, the presence of Lateral hinge teeth which develop after
metamorphosis are unique and consistent criteria for distinguishing juvenile mussels of the
genus Perna. The development and fate of provincular, primary lateral, secondary lateral
and dysodont hinge teeth is presented in a series of scanning electron micrographs. Dis-
tinguishing features of soft-part anatomy and adult shells are summarized. Several anatomical
features are discussed with respect to a possible trend in specialization among the genera
Perna, Choromyfilus, Aulacomya and Mytilus.

The Mytilidae, or true mussels, demonstrate a great deal of variation in mor-

phological features which are taxonomically important in the Bivalvia. Thus the
taxonomic status of species within the Mytilidae is often confused. For want of
reliable morphological features by which to distinguish species and genera, the
inconsistent or “plastic” character of gross adult shell morphology has, in the
past, shaped the family’s hierarchy. The existence of physiological races and a
wide range of eco-morphs in the Mytilidae has complicated the interpretation of
experimental evidence and created a taxonomic challenge for the researcher
working with mytilids. Only recently have warm water mussels of the genus
Perna been studied extensively, primarily because of their value as a food
resource. Nomenclatural errors have persisted in these studies which have
separately treated synonymous species of mussels and reported on eco-physio-
logical differences between two or more races of the same species. In this paper,
I describe several larval and adult characteristics of the three living species cur-
rently placed in the genus Perna (P. perna, P. viridis and P. canuliculus), briefly
review the historical development and geographical distribution of the species,
and present criteria for distinguishing larvae and adults of the genus Perna from
those of the genus Myths.

HISTORICAL REVIEW
In 1932, the rosters of Index Animalium reported on the use of approximately
400 different species names in the genus’ Mytilus and another 71 names in the
genus Perna. Unfortunately, the name Perna had been used to describe two
genera of mytilids, Perna (Retzius, 1788) and Modiolus (H. and A. Adams, ISSS),
and a genus of Pteriacea, Zsognomon (Bruguiere, 1792). When Retzius (1788) set
forth the genus Perna, he listed the type specimen as Perna magellanica which
according to Lamy (1936-1937) is synonymous with Linnaeus’ Mya perna and
whose holotype was presumably collected from the Straits of Magellan. Both are
properly referred to as Perna perna (L.) (Lamy, 1936-1937). P. magellanica as
described by Retzius must not be confused with Mytilus magellanicus Chemnitz
which, as Soot-Ryen (1955) details, probably belongs to the genus Aulacomya.
Linnaeus (1758) first described Mytilus viridis while Gmelin (1791) is credited
for M. cunaliculus in spite of Martyn’s (1784) earlier work which has been de-
858
 SIDDALL: CLARIFICATION OF THE GENUS PERNA 859

termined to possess no status in zoological nomenclature (I.C.Z.N. opinion 456).

The several works of Born, Chemnitz, Dillwyn, Gmelin and Lamarck expanded
the list of species in the genus Perna but Hanley (1843, 1855) temporarily reversed
this trend by lumping together several species. One notable point of confusion
was Lamarck’s use of M. latus (a synonym for Choromytilus chorus; Soot-Ryen,
1952), a name also employed by Chemnitz for the New Zealand green mussel,
now referred to as Perna canaficulus (Fleming, 1959). In Hanley’s work (1843)
the synonymy of M. viridis L. and M. smurugdinus Chemnitz was discussed as
was the synonymy between M. cunuliculus (Gmelin) and M. lutus Chemnitz.
March (1853) set aside the genus Chloromyu for those species belonging to Ret-
zius’ earlier genus, Pernu. The use of the genus Chloromyu added to the con-
fusion, persisting beyond Lamy’s later work.
Following the turn of the century, von Ihering (1901, 1907) and particularly
Jukes-Browne (1905) discussed hinge and ligament structures and muscle scars
as bases for establishing the taxonomic hierarchy of the Mytilidae. This work laid
the foundation for Lamy’s (1936-1937) analysis of museum specimens involving
both the genera Mytilus and Pernu. The confused taxonomic interrelationships
involving Mytilus and Pernu which developed in the 19th century were simplified
by Lamy’s comprehensive works followed by those of Soot-Ryen (1952, 1955).
Dodge (1952) still regarded Chloromyu as a subgenus of Mytilus. In 1952, Soot-
Ryen divided the genus Pernu (at the time, still referred to as Chloromyu) into
two groups: 1) those having a pitted resilial ridge belonging to the genus Chlo-
romyu, and 2) those having a compact resilial ridge forming a new genus, Cho-
romytilus with the genotype C. chorus Molina 1782. Soot-Ryen (1955) showed
Chloromyu to be an invalid synonym for Pernu and clarified much of the taxo-
nomic nomenclature of the mytilids by retaining Pernu (Retzius) 1788 for those
species of mytilids resembling Mytilus s. str. but which have a pitted resilial ridge
and discontinuous posterior retractor muscle scars and lack an anterior adductor
muscle. Dance (1974) placed C. chorus Molina in the genus Pernu but gave no
new evidence supporting this change. The close taxonomic relationships among
the genera Mytilus, Pernu, Choromytilus, and Aulucomyu are briefly described
by Soot-Ryen (1952). In Beauperthuy’s (1967) analysis of the Venezuelan mytil-
ids, including P. perna, the genus “Chloromytilus” appears most probably as a
typographical error referring to Soot-Ryen’s valid genus Choromytilus.
Table 1 is not a comprehensive survey of the historical development of the
genus Pernu, but it summarizes a great many synonymies involving the three
species.

MATERIALSAND METHODS
Larvae and adults of the three species of Pemu were collected from 21 locations distributed
throughout known geographical ranges including type localities. Specimens of P. perna were collected
in Venezuela, Brazil and South Africa, P. vin’dis from Pakistan, India (both coasts), Singapore and
the Philippines (two stations 600 km distant), and P. canaliculus from the north island of New
Zealand. Observations of soft-part anatomy were made from adult specimens preserved in buffered
5% formalin. Live adults of P. pevna (Cumana, Venezuela) and P. viridis (Bacoor Bay, Philippines)
were imported to Miami and quarantined. Current New Zealand governmental policy does not permit
exportation of living specimens of P. cana/iculu~s.
Live P. perna and P. viridis adults were held in running seawater (28-32c/r0 at 22-26°C) while being
conditioned for spawning. Adults and developing larvae were fed a mixed phytoplankton diet of
Chaetoceros calcitrum, Dunaliellu tertiolecta, Isochrysis gulbana, Monochtysis lutheri, and Tetra-
selmis suecicn. Ripe adults spawned regularly within one half hour of being exposed to 35°C seawater
for 10 min. Details of the culture procedures are outlined in Siddall (1979a). Larval cultures were
maintained for 24 days while samples of larvae were withdrawn and preserved in Carriker’s (1950)
solution every 24 h for later examination. The lack of suitable substrates for settlement within the
860 BULLETIN OF MARINE SCIENCE, VOL. 30, NO. 4, 1980

Table I. Important works describing P. perna, P. viridis and P. canaliculus and summary of syn-
onymies

Perna pernu (Linnaeus) 1758

1758 Myu perna Linnaeus, Syst. Nat., Ed. X
1780 Mytilus pictus Born, Test. Mus. Caes. Vindob.
1785 Mytilus hfricanus Chemnitz, Conch. Cab. VIII
1791 Mytilus afer Gmelin, Svst. Nat., Ed. XIII
1817 Mytilus elongatus Lamarck <non Chemn.), Anim. s. Vert. VI
1819 Mytilus perna L. Lamarck, Anim. s. Vert. VI
1843 Mytilus perna L. Hanley, Cat. Rec. Biv. Sh.
1855 Mya perna L. Hanley, Ipsa Linn. Conch.
1901 Mytilus perna L. von Ihering, Proc. Malac. Sot. Lond. IV
1905 Mytilus (Chloromya) perna L. Jukes-Browne, Proc. Malac. Sot. Lond. VI
1905 Mytilus (Chloromya) pictus Born Jukes-Browne, Proc. Malac. Sot. Lond. VI
1905 Mytilus (Chloromya) afer Gmelin Jukes-Browne, Proc. Malac. Sot. Lond. VI
1907 Mytilus perna L. von Ihering, Anal. Mus. Nat. Buenos Aires XIV
1919 Mytilus (Chloromya) perna L. Lamy, Bull. Mus. Hist. Nat. XXV
1931 Mytilus (Chloromya) perna L. Lamy, Bull. Mus. Hist. Nat. III. 2e Ser.
1936 Mytilus (Chloromyu) perna L. Lamy, Rev. Mytilidae J. Conchyl. LXXX
1952 Chloromyu perna L. Dodge, Bull. Amer. Mus. Nat. Hist., 100
1955 Perna perna L. Soot-Ryen, Allan Hancock Pac. Exped. 20(l)
1964 Perna perna L. Barnard, Ann. S. Afr. Mus. 47
1965 Mytilus (Chloromya) venezolanus Andreu, Inst. Invest. Pesq. Barcelona 5
1965 Chloromya perna L. Padilla, Rev. Inst. Invest. Pesq. Uruguay
l(4)
1967 Perna perna L. Beauperthuy, Bol. Inst. Oceanogr. Univ.
Oriente, VI
1969 Perna perna L. Nordsieck, Die Europaischen Meeresmuscheln
Perna picta picta Born Nordsieck, Die Europaischen Meeresmuscheln
1973 Pernu perna L. Lubet, F.A.O. Synopses sur les Peches,
Vol. 88
1973 Perna perna L. Zaoualt, Rapp. Comm. Inst. Mer. Med. 22
1974 Perna perna L. Day, Guide Mar. Life. S. Afr. Shores
1975 Pernu perna L. Rios, Brazilian Mar. Mall. Iconography
1976 Perna (Perna) picta (Born) Buccheri and Palisano, Conchiglie 12

Pernu viridis (Linnaeus) 1758

1758 Mytilus viridis Linnaeus, Syst. Nat., Ed. X
1785 Mytilus smaragdinus Chemnitz. Conch. Cab. VIII
1791 Mytilus smaragdinus Chemnitz Gmelin, Syst. Nat., Ed. XIII
1819 Mytilus opalus Lamarck, Anim. s. Vert. VI
1819 Mytilus smaragdinus Gmelin Lamarck, Anim. s. Vert. VI
1843 Mytilus smarngdinus Hanley, Cat. Rec. Biv. Sh.
1843 Mytilus viridis L. Hanley, Cat. Rec. Biv. Sh.
1855 Mytilus viridis L. Hanley, Ipsa Linn. Conch.
1857 Mytilus smaragdinus Chemnitz Reeve, Conch. Icon.
1905 M. (Chloromya) smaragdinus Jukes-Browne, Proc. Malac. Sot.
Chemnitz Lond. VI
1906 Mytilus viridis L. Dautzenberg and Fischer, J. Conchyl. LIV
1936 Mytilus (Chloromya) viridis L. Lamy, Rev. Mytilidae, J. Conchyl. LXXX
1937 Mytilus smaragdinus Chemnitz Serene, Invent. Invert. Mar. Indochine
1937 Mytilus viridis L. Serene, Invent. Invert. Mar. Indochine
1950 Mytilus smaragdinus Chemnitz Suvatti, Fauna Thailand
1950 Mytilus viridis L. Suvatti, Fauna Thailand
1952 Chloromya viridis L. Dodge, Amer. Mus. Nat. Hist., 100
1968 Mytilus viridis L. Cheriyan, Symp. Mollusca 3
1974 Perna viridis L. Ahmed, Bol. Inst. Oceanogr. Univ.
Oriente XIII
1974 Perna viridis L. Dance, Encyc. of Shells
 SIDDALL: CLARIFICATION OF THE GENUS PERNA 861

Table 1. Continued

Perna canalicu/us (Gmelin) 1791

1784 Mytilus canuliculus Martyn, Univ. Conch. II
178.5 Mytilus lutus Chemnitz (non Lamarck), Conch. Cab. VII
1791 Mytilus cunaliculus Gmelin, Syst. Nat., Ed. XIII
1843 Mytilus cunuliculus Martyn -Hanley, Cat. Rec. Biv. Sh.
1873 Mytilus lutus Chemnitz Hutton (non Lamarck), Cat. Mar. Moll.
1873 Mytilus smarugdinus Hutton (non Chemnitz), Cat. Mar. Mall.
1913 M. (Chloromyu) cunuliculus Suter, Man. N. Zealand Mall.
Martyn
1924 Mytilus cunuliculus Martyn Odhner, Vidensk, Medd. fra Dansk, Foren.,
Bd. 77
1959 Pernu cunuliculus Martyn Fleming, Trans. Roy. Sot. N. Zealand 87

cultures forced a delay in completion of metamorphosis. Onset of metamorphosis was determined by

presenting a suitable substrate to a subsample of larvae; if more than half of the larvae secreted a
byssus, that culture was deemed capable of initiating metamorphosis. This procedure was carried out
every 12 h beginning 6 days after fertilization. These metamorphosing pediveligers were cultured
separately through completion of metamorphosis (secretion of the dissoconch shell; see Bayne, 1965).
Preserved samples were cleaned of organic matter in a buffered l-2% sodium hypochlorite solution
for 30 set to 3 min, rinsed, air-dried and then examined with an AMR-900 scanning electron micro-
scope. Structures associated with the hinge line, i.e., provincular and lateral hinge teeth and ligament
pit, were examined in detail. Micrographs of periostracum cross sections were made from fractures
of juvenile (0.5 cm) and small adult (l-2 cm) shells.

RESULTS
Morphogenesis of the hinge lines in larvae and juveniles of the three species
is shown in Figure 1. At 26”C, the prodissoconch I stage (“D” stage larvae) is
reached 14-18 h after fertilization by larvae of P. perna and P. viridis. At the
prodissoconch II stage, larval shells of all three species have a variable number
of provincular hinge teeth (Fig. lA-C). Prior to metamorphosis (upper two rows
of Fig. l), the ends of the provinculum (as defined by Rees, 1950) widen as the
outer hinge teeth enlarge. Throughout larval growth, the number of provincular
hinge teeth increases.
At 26”C, both P. perna and P. viridis secrete the first byssus lo-20 days after
fertilization. This indicates the onset of metamorphosis. In pediveligers preserved
at this time, the ligament pit is apparent just below the central area of the pro-
vinculum (li, Fig. 1G). Also a series of what will be referred to as primary lateral
hinge teeth (1,) Fig. 1H) forms on the dorsal margin posterior and adjacent to the
provincular teeth of both right and left valves. These primary lateral teeth, like
the provincular teeth, interdigitate with those of the opposing valve. Between 10
and 18 primary lateral teeth are present in pediveligers which are allowed to
complete metamorphosis (Fig. lJ-L). In contrast, metamorphosed postlarvae of
Mytihs edulis do not develop primary lateral hinge teeth immediately adjacent
to the provincular teeth (Cox, 1969).
However, following completion of metamorphosis, plantigrades of Perna (and
Mytilus; Cox, 1969 and Le Pennec and Masson, 1976) develop two additional
series of teeth. A series of hinge teeth (1, in Fig. 2) appears on the dorsal shell
margin immediately posterior and adjacent to the primary lateral hinge teeth.
These hinge teeth, herein referred to as secondary lateral teeth, are distinctly
larger than the primary lateral teeth in Perna. The differences between primary
and secondary lateral teeth are most clearly shown in the upper plate of Figure
X62 RUI I E 1IN OF MARINF SCIF,N(‘F. VOI 70. NO 4. 1980

R perna I I I? viridis I I R canaliculus

I 1

Figure 1. Morphogenesis of hinge lines in three species of Pcmtr; all views are of left valves with
the exception of Ik and II. a-c, early prodissoconch II shells (1000x): shell lengths are: a = IO2
wrn, b = 105 Wm. c = 110 pm. d-f, late prodissoconch II shells, 6 to 8 days old (500x); shell lengths
are: d = I98 pm, e = 190 pm, f = 180 +m. g-i, metamorphosing pediveligers, 10 to I2 days old
(500x): shell lengths are: g = 230 wrn, h = 215 Frn, i = 235 pm. j-l, plantigrades, 18 to 24 days
old (250x); maximum shell dimensions are: j = 315 pm, k = 310 pm. I = 2X5 Wm. m-o, juvenile
mussels, 40 to 45 days old (50x); maximum shell dimensions are: m = 500 pm, n = 430 pm, o = 480
Wm. dy = dysodont teeth, I, = primary lateral hinge teeth, I, = secondary lateral hinge teeth. Ii =
ligament pit, and p 7 provincular hinge teeth.

2. While provincular teeth are oriented perpendicularly to the provinculum, the

secondary laterals are oriented longitudinally and more nearly parallel to the
adjacent shell margin. In metamorphosed Mytilus gallr,provin~icr/i,v, Le Pennec
and Masson (1976) described these posterior laterals as having the same longi-
tudinal or fan-shaped orientation. In juvenile Porno, the definitive ligament ap-
pears along the posterodorsal shell margin and subsequently expands obscuring
the primary and secondary lateral hinge teeth.
The second series of teeth develops anterior to the beaks and somewhat distant
from the provincular teeth (dy in Fig. 2). These anteroventral teeth are not part
of the dorsal hinge structure, are not covered by the ligament and are the only
hinge teeth which remain distinctly visible in the adult. As discussed by Cox
(1969), these are the typical dysodont of mytilids. Le Pennec and Masson (1976)
refer to these dysodont teeth as ventral cardinal teeth in juvenile Mytilus gullo-
provinc’icrlis. In Pcrna, dysodont teeth are oriented perpendicularly to the an-
teroventral shell margin in juveniles, yet oriented longitudinally in adults. In
members of the genus Pcrtztl, dysodont teeth usually number two (Fig. 2, lower
plate), interdigitate with one or two teeth present on the opposing valve, and may
be formed by the radial ridges of the lunule (Soot-Ryen, 1955). The interspecific
differences between P. pcrnn and P. viridis regarding the position and number
of dysodont teeth as mentioned by Dodge (1952) were not consistent in the ma-
terial examined in this study. As Jukes-Browne (1905), Lamy (1936), Beauperthuy
(1967) and others have observed, the number of dysodont teeth (l-2) and position
(right or left valve) vary.
 \IDI)Al I. Cl AKIFICAIION OF IHF GFNIIS PhHNA 863

Figure 2. (upper) High magnification detail of primary (1,) and secondary (I,) lateral hinge teeth of
left valve of a juvenile P. c-~rntr/ic~u/u,r (maximum shell dimension = 480 pm). (lower) Dysodont hinge
teeth (dy) along anteroventral shell margin (left valve) of juvenile P. p~~‘n~tr (maximum shell dimen-
sion = 3 mm).

Though the number and size of the provincular, primary lateral and secondary
lateral teeth show considerable variation, there are no significant differences
among the three species of Pernu studied. Examination of preserved larval and
adult specimens of all three species confirmed Seed’s (1968) contention that shell
X64 IllI IXTIN OF MARINE S(‘IRN(‘I-‘., VOI.. 30, NO. 4, 19X0

cana liculus
Figure 3. Ultrastructural details of three species of Pcvnrr: a-c, provincular hinge teeth showing
ridges along teeth CWOOx); d-f. primary lateral hinge teeth of dorsal shell margins demonstrating
coarse ultrastructure (2500x): g-l, cross sections of three-layered periostracum as seen in juvenile
shell fractures (500x).

shape and thickness are characters of little taxonomic value in the Mytilidae.
Variations in shell coloration and patterns are considerable in all material; how-
ever, light colored zigzag markings are most common in young P. cmncrliculm
specimens. P, ~~rrz~ adults are typically brown to red-maroon with irregular areas
of light brown and green. Brilliant green and blue-green predominate in P. viridis
juveniles while adult shells are less brilliant and have a greater proportion of
brown. In older specimens of all species, abrasion of the external anterior surface
removes the periostracum exposing the white to pink outer shell surface.
Interspecific variations in soft-part anatomy are inconsistent and difficult to
document. Bifurcating papillae or tentacles along the mantle edge are most pro-
nounced in P. perncr, less co in P. I*iridi.s. The upper edges of the gill lamellae
are more strongly attached to the mantle in all three species of Perntr than in
Mytilrrs eclrrlis. Finally, in P. pcrnrr and P. Lliriciis, two separate gonoductc lead
to the mantle cavity; distinct streams of gametes are visible in spawning adults.
This is in contrast to M. cdulis which has a single gonoduct opening (Field, 1922).
Figure 3 presents ultrastructural details of the hinge and periostracum. Figures
3A-C reveal the ridged nature of the provincular hinge teeth. Where right and
left valves remained articulated during examination, it was apparent that these
ridges consistently interlock with their counterparts on the teeth of the opposing
 SIDDALL: CLARIFICATION OF THE GENUS PERNA 865

R canaliculus Mytilus edulis

Figure 4. Muscle scar patterns on cleaned right valves of Prmcr and Myfilus. Contrast two-part
retractor muscle scars of Prrnu with the single scar of Myfilus. Anterior adductor muscle of Mytilus
is only partially visible in the anteroventral angle of the shell. No anterior adductor is present in the
genus Perna. Drawn l/2 life size.

valve. Lutz and Hidu (1979) also observed these ridges in other mytilids. Pre-
sumably the function of the ridges is to stabilize motions of larval valves which
must gape widely to accommodate the relatively large velum (Stanley, 1978, and
Wailer’s discussion following Stanley’s paper). Stanley (1978) refers to these
ridges in the Trigoniidae as secondary dentition, a term which should not be
confused with secondary lateral teeth defined above. Figures 3D-F depict the
ultrastructure of the primary lateral teeth which contrasts with the smooth or
amorphous character of the provincular teeth. Because this feature is consistent
and restricted to the primary lateral teeth, it is not an artifact resulting from the
brief exposure to hypochlorite during sample preparation. Carriker (1979)
describes in detail the minor ultrastructural effects of cleaning molluscan shells
with sodium hypochlorite. Figures 3g-i show the three layered structure of the
juvenile periostracum resembling that described by Dunachie (1963) for M. edulis.
Analysis of X-ray dispersion during scanning electron microscope examination
confirmed the organic nature of the periostracum but also revealed an elevated
calcium content in the middle layer 2.5 times greater than that of the surrounding
layers of periostracum. Such levels of calcium are probably related to the pres-
ence of calcified, spicule-like structures in the periostracum (Carter and Aller,
1975). Though the coloration of the adult mytilid is a feature of the periostracum,
the presence or distribution of trace elements within the periostracum did not
correlate with adult color patterns.
Variations in the pattern of muscle scars left on juvenile and adult shells are
shown in Figure 4. On a macroscopic level, the shell layer associated with sites
of muscle attachment, the myostracum, may be seen as muscle scars and the
pallial line. The discontinuous nature of the retractor muscle scar is one of the
definitive characters of the genus Perna. The anterior-most component of the
retractor muscle complex is attached to the shell at a point removed from the
866 BULLETIN OF MARINE SCIENCE.VOL. 30. N0.4. 1980

1.’
\w 3 E perna
Bvm
B canaliculus

Figure 5. Geographical distribution of the three species of Prmrr.

point of attachment of posterior retractors. This results in a two-part or discon-

tinuous muscle scar. In contrast, the middle and posterior retractor muscles in
Mytilus are united and leave a continuous band of myostracum along the dorsal
margin of the pallial line (see also Fig. 4). In Figure 4, the white resilial ridge,
which is distinctly pitted or porous in both Perna and Mytilus, appears as the
stippled band along the dorsoanterior shell margins. None of the three species of
Perna have an anterior adductor muscle. Mytilus edulis, on the other hand, does
have an anterior adductor muscle, although relatively small, which attaches along
the anteroventral margin (partially seen in Fig. 4). Newell (1969) stated that an
anterior adductor is present in P. perna juveniles but such was not the case for
material examined in this study.

DISCUSSION
Geographic distributions of the species of Perna are shown in Figure 5. This
figure was compiled primarily from the literature with minor geographical exten-
sions resulting from personal communications with the many researchers and
malacologists who assisted in the collection of material for this study. Reports in
the literature of P. perna in cooler waters south of Rio de la Plata, Argentina, to
the Straits of Magellan are open to question (Penchaszadeh, personal communi-
cation). Beauperthuy (1967, p. 35) discusses the possibility that the type locality
of P. perna, the Straits of Magellan, was erroneously assigned. Although Lamy
(1920, 1936-1937) and Soot-Ryen (1955) do not rule out the presence of a mussel
conspecific with P. perna in South America (M. achatinus, M. elongatus?), the
preserved material examined in this study indicated a single species, P. perna,
present in Venezuela, along the coast of Brazil at Recife, in the Straits of Ma-
gellan, and along the African continent. Experimental evidence on the limited
duration of planktonic stages in Perna (Siddall, 1979 and unpublished data) in-
dicates that this distribution could not be a result of long distance transport of P.
perna across the North or South Atlantic oceans. The possibility for widespread
 SIDDALL: CLARIFICATION OF THE GENUS PERNA 867

Table 2. Summary of distinguishing characteristics of four genera in the Mytilidae

Mpfr/r,.v A!ll~KO?7ly~l Choromyrilus PfWn

Anterior adductor Anterior adductor No anterior No anterior adductor

always present present in young; adductor
absent in adult
One retractor scar One retractor scar One retractor scar, Two retractor scars
sometimes two
No primary lateral Condition unknown Condition unknown 10 to 18 primary
teeth lateral teeth
One to two Ridge, no teeth One to two Usually two dysodont
dysodont teeth dysodont teeth teeth, sometimes
one
Pitted resilial ridge Compact or dense Compact or dense Pitted resilial ridge
resilial ridge resilial ridge
Smooth shell Strongly ribbed Faintly ribbed shell Smooth shell
shell

geographical distribution by step-wise larval dispersal (“island hopping”) does

exist for P. viridis in the Indo-Pacific.
Nordsieck (1969) refers to a near-cosmopolitan distribution for P. perna but
he also differentiates two other mussels, P. picta picta (an eco-morph of P. perna
from the Mediterranean and North Africa) and P. picta mau-
retcrnica (= M. gnlloprovincialis: Lamy, 1936). Buccheri and Palisano (1976) also
argue for the presence of P. picta (Born) as a separate species in the Mediter-
ranean. Beauperthuy’s (1967) statement that P. perna extends to New Zealand
and the tropical Pacific is unsubstantiated and probably reflects the great mor-
phological variation within the genus Perna. Because of the degree of variation
in characters of taxonomic importance within the genus Perna, it is difficult to
reliably distinguish the species without knowing from what locality the material
was collected. In this respect, it is fortunate that the geographical ranges of the
three species are not known to overlap. Aside from the minor differences in
coloration already noted, only the presence of enlarged sensory papillae along
the mantle margins in P. perna aids in species identification.
At the generic level, however, there are a variety of distinguishing character-
istics which are summarized in Table 2. In the material examined for this work,
the presence or absence of anterior adductor muscles and variations in the number
of retractor muscle scars are consistent characters of both juveniles and adults
on which to base taxonomic differentiation of Perna and Mytilus. The order of
the genera listed in Table 2 (left to right)‘is based on a trend involving these two
anatomical variations. The anterior adductor is always present in Mytifus, absent
only in older Aulacomya specimens, and completely absent in Choromytilus and
Perna. There is a single retractor muscle scar in Mytifus and Aufacomya, one
and occasionally two in Choromytilus, and two in Perna. Until the presence or
absence of primary lateral teeth in Aulacomya and Choromytilus can be deter-
mined, this criterion neither supports nor refutes any trend suggested by muscle
arrangements. However, the order of the trend (Mytilus-Aulacomya-Choromy-
tilus-Perna) does break down when we consider dysodont hinge teeth, resilial
ridges, and external shell sculpture. At this point, there is no reason to emphasize
the taxonomic value of any one set of these characters over another. Certainly
more information is required before this trend may be properly evaluated. The
868 BULLETIN OF MARINE SCIENCE, VOL. 30, NO. 4, 1980

underlying significance of the suggested trend, if there is any, remains unclear

but may relate to the degree of evolutionary “specialization” of each genus. From
that standpoint, Perna, having lost the anterior adductor muscle, divided the
retractor muscle complex, and developed primary lateral hinge teeth and branch-
ing papillae on the mantle margin may be more specialized, yet the advantages
of such morphological alterations are not obvious. Some support for this trend
in specialization may be found in Ahmed (1974) and Ahmed and Sparks (1970),
who showed that the diploid chromosome number in M. e&/is, M. californianus,
and P. perna is 28 while in P. viridis it is 30. Based on the potential mechanisms
for such numerical differences in karyotypes, Ahmed (1974) tentatively concluded
that the genus Perna is more specialized than Mytilus and that P. viridis is more
specialized than P. perna.
In his 1969 review, Cox points out that following metamorphosis, M. edulis
develops hinge teeth “along the posterodorsal margin some distance beyond the
row of crenulations persisting from the prodissoconch, which are obliterated as
the ligament extends posteriorly. Similar teeth appear along the margin anterior
to the beak, also beyond the row of crenulations, and some of these persist to
form the dysodont teeth of the adult mussel.” I have termed Cox’s posterodorsal
series of teeth secondary laterals. Primary lateral teeth immediately adjacent to
the provincular teeth are not mentioned by Cox or by Lutz and Hidu (1979) who
examined larvae and early post-larvae of M. edulis or by Le Pennec and Masson
(1976) working with M. galloprovincialis. In Perna, both the primary lateral teeth
and the ligament pit are present in stage 3 pediveligers (Bayne, 1965) which had
not yet completed metamorphosis. Secondary lateral and dysodont teeth appear
very soon after (less than 24 h) completion of metamorphosis as indicated by
secretion of the dissoconch shell. However, because of the 24-h sampling interval
used in this study, it is not possible to further resolve the timing of these events
in Perna. Furthermore, one would expect significant differences in the timing of
ontogenetic events between Mytilus, a temperate genus, and the subtropical to
tropical species of Perna. Though larval hinge structure appears to be a relatively
conservative taxonomic character (Rees, 1950; Cox, 1969), there are significant
variations within the families Ostreidae (Ranson, 1948) and Mytilidae (present
study).
The presence of primary lateral hinge teeth allows us to differentiate
larvae of Perna from those of Mytilus. Adults of the genera are most reliably
distinguished by patterns of muscle scars. Fewer criteria are available for distin-
guishing among the three species of Perna ; geographic origin, coloration and
some aspects of soft-part morphology are useful. Recognition of environmental
variations which favor the development of ecomorphs of these mussels should
minimize the current tendency to confuse the taxonomic status and nomenclature
of the three species of Perna.

ACKNOWLEDGMENTS
Special thanks are due Drs. T. R. Wailer, R. A. Lutz and E. S. Iversen for critical review of the
manuscript. I am also indebted to the great number of researchers and malacologists who collected
and forwarded preserved material used in this effort. This is a contribution from the Rosenstiel School
of Marine and Atmospheric Science, University of Miami.

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 SIDDALL: CLARIFICATION OF THE GENUS PERNA 869

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DATE ACCEPTED: October 10, 1979.

ADDRESS: Rosenstiel School of Marine rend Atmospheric Science, 4600 Rickenhrrcker Ctrrrsr~\~ry.
Miami, Florida 33149.